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Yeasts are
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
, single-celled
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s classified as members of the
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately fr ...
kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
are currently recognized. They are estimated to constitute 1% of all described fungal species. Yeasts are unicellular organisms that evolved from
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially ...
ancestors, with some species having the ability to develop multicellular characteristics by forming strings of connected budding cells known as
pseudohyphae A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one o ...
or false hyphae. Yeast sizes vary greatly, depending on species and environment, typically measuring 3–4 
µm The micrometre ( international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer ( American spelling), also commonly known as a micron, is a unit of length in the International System of Uni ...
in
diameter In geometry, a diameter of a circle is any straight line segment that passes through the center of the circle and whose endpoints lie on the circle. It can also be defined as the longest chord of the circle. Both definitions are also valid f ...
, although some yeasts can grow to 40 µm in size. Most yeasts reproduce asexually by mitosis, and many do so by the asymmetric division process known as
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
. With their single-celled growth habit, yeasts can be contrasted with molds, which grow
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
e. Fungal species that can take both forms (depending on temperature or other conditions) are called
dimorphic fungi Dimorphic fungi are fungi that can exist in the form of both mold and yeast. This is usually brought about by change in temperature and the fungi are also described as thermally dimorphic fungi. An example is '' Talaromyces marneffei'', a huma ...
. The yeast species ''
Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have b ...
'' converts
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may o ...
s to
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
and alcohols through the process of fermentation. The products of this reaction have been used in baking and the production of alcoholic beverages for thousands of years. ''S. cerevisiae'' is also an important model organism in modern cell biology research, and is one of the most thoroughly studied eukaryotic microorganisms. Researchers have cultured it in order to understand the biology of the eukaryotic cell and ultimately human biology in great detail. Other species of yeasts, such as '' Candida albicans'', are opportunistic pathogens and can cause
infections An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable di ...
in humans. Yeasts have recently been used to generate electricity in microbial fuel cells and to produce
ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a ...
for the
biofuel Biofuel is a fuel that is produced over a short time span from biomass, rather than by the very slow natural processes involved in the formation of fossil fuels, such as oil. According to the United States Energy Information Administration ...
industry. Yeasts do not form a single taxonomic or
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
grouping. The term "yeast" is often taken as a synonym for ''Saccharomyces cerevisiae'', but the phylogenetic diversity of yeasts is shown by their placement in two separate
phyla Phyla, the plural of ''phylum'', may refer to: * Phylum, a biological taxon between Kingdom and Class * by analogy, in linguistics, a large division of possibly related languages, or a major language family which is not subordinate to another Phy ...
: the Ascomycota and the Basidiomycota. The budding yeasts or "true yeasts" are classified in the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
Saccharomycetales, within the phylum Ascomycota.


History

The word "yeast" comes from
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlers in the mid-5th ...
''gist'', ''gyst'', and from the
Indo-European The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, Du ...
root ''yes-'', meaning "boil", "foam", or "bubble". Yeast microbes are probably one of the earliest domesticated organisms. Archaeologists digging in Egyptian ruins found early grinding stones and baking chambers for yeast-raised bread, as well as drawings of 4,000-year-old bakeries and breweries. Vessels studied from several archaeological sites in Israel (dating to around 5,000, 3,000 and 2,500 years ago), which were believed to have contained alcoholic beverages (beer and mead), were found to contain yeast colonies that had survived over the millennia, providing the first direct biological evidence of yeast use in early cultures. In 1680, Dutch naturalist Anton van Leeuwenhoek first microscopically observed yeast, but at the time did not consider them to be
living organisms In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fun ...
, but rather globular structures as researchers were doubtful whether yeasts were algae or fungi. Theodor Schwann recognized them as fungi in 1837. In 1857, French microbiologist Louis Pasteur showed that by bubbling oxygen into the yeast broth,
cell growth Cell growth refers to an increase in the total mass of a cell, including both cytoplasmic, nuclear and organelle volume. Cell growth occurs when the overall rate of cellular biosynthesis (production of biomolecules or anabolism) is greater th ...
could be increased, but fermentation was inhibited – an observation later called the " Pasteur effect". In the paper "''Mémoire sur la fermentation alcoolique,''" Pasteur proved that alcoholic fermentation was conducted by living yeasts and not by a chemical catalyst. By the late 18th century two yeast strains used in brewing had been identified: ''Saccharomyces cerevisiae'' (top-fermenting yeast) and '' S. carlsbergensis'' (bottom-fermenting yeast). ''S. cerevisiae'' has been sold commercially by the Dutch for bread-making since 1780; while, around 1800, the Germans started producing ''S. cerevisiae'' in the form of cream. In 1825, a method was developed to remove the liquid so the yeast could be prepared as solid blocks. The industrial production of yeast blocks was enhanced by the introduction of the filter press in 1867. In 1872, Baron Max de Springer developed a manufacturing process to create granulated yeast, a technique that was used until the first World War. In the United States, naturally occurring airborne yeasts were used almost exclusively until commercial yeast was marketed at the Centennial Exposition in 1876 in Philadelphia, where Charles L. Fleischmann exhibited the product and a process to use it, as well as serving the resultant baked bread. The mechanical refrigerator (first patented in the 1850s in Europe) liberated brewers and winemakers from seasonal constraints for the first time and allowed them to exit cellars and other earthen environments. For John Molson, who made his livelihood in
Montreal Montreal ( ; officially Montréal, ) is the second-most populous city in Canada and most populous city in the Canadian province of Quebec. Founded in 1642 as '' Ville-Marie'', or "City of Mary", it is named after Mount Royal, the triple- ...
prior to the development of the fridge, the brewing season lasted from September through to May. The same seasonal restrictions formerly governed the distiller's art.


Nutrition and growth

Yeasts are
chemoorganotroph Primary nutritional groups are groups of organisms, divided in relation to the nutrition mode according to the sources of energy and carbon, needed for living, growth and reproduction. The sources of energy can be light or chemical compounds; the ...
s, as they use
organic compound In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. Th ...
s as a source of energy and do not require sunlight to grow. Carbon is obtained mostly from hexose sugars, such as
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...
and fructose, or disaccharides such as sucrose and maltose. Some species can metabolize
pentose In chemistry, a pentose is a monosaccharide (simple sugar) with five carbon atoms. The chemical formula of many pentoses is , and their molecular weight is 150.13 g/mol.organic acid An organic acid is an organic compound with acidic properties. The most common organic acids are the carboxylic acids, whose acidity is associated with their carboxyl group –COOH. Sulfonic acids, containing the group –SO2OH, are ...
s. Yeast species either require oxygen for aerobic
cellular respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
(
obligate aerobe An obligate aerobe is an organism that requires oxygen to grow. Through cellular respiration, these organisms use oxygen to metabolise substances, like sugars or fats, to obtain energy. In this type of respiration, oxygen serves as the terminal ...
s) or are anaerobic, but also have aerobic methods of energy production (
facultative anaerobe A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are '' Staphylococ ...
s). Unlike
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
, no known yeast species grow only anaerobically ( obligate anaerobes). Most yeasts grow best in a neutral or slightly acidic pH environment. Yeasts vary in regard to the temperature range in which they grow best. For example, '' Leucosporidium frigidum'' grows at , '' Saccharomyces telluris'' at , and '' Candida slooffi'' at . The cells can survive freezing under certain conditions, with viability decreasing over time. In general, yeasts are grown in the laboratory on solid growth media or in liquid broths. Common media used for the cultivation of yeasts include
potato dextrose agar Potato dextrose agar (BAM Media M127
from the U.S.
or
potato dextrose broth Potato dextrose agar (BAM Media M127
from the U.S.
, Wallerstein Laboratories nutrient
agar Agar ( or ), or agar-agar, is a jelly-like substance consisting of polysaccharides obtained from the cell walls of some species of red algae, primarily from ogonori (''Gracilaria'') and "tengusa" (''Gelidiaceae''). As found in nature, agar i ...
, yeast peptone dextrose agar, and yeast mould agar or broth. Home brewers who cultivate yeast frequently use dried malt extract and agar as a solid growth medium. The fungicide cycloheximide is sometimes added to yeast growth media to inhibit the growth of ''
Saccharomyces ''Saccharomyces'' is a genus of fungi that includes many species of yeasts. ''Saccharomyces'' is from Greek σάκχαρον (sugar) and μύκης (fungus) and means ''sugar fungus''. Many members of this genus are considered very important in f ...
'' yeasts and select for wild/indigenous yeast species. This will change the yeast process. The appearance of a white, thready yeast, commonly known as kahm yeast, is often a byproduct of the lactofermentation (or pickling) of certain vegetables. It is usually the result of exposure to air. Although harmless, it can give pickled vegetables a bad flavor and must be removed regularly during fermentation.


Ecology

Yeasts are very common in the environment, and are often isolated from sugar-rich materials. Examples include naturally occurring yeasts on the skins of fruits and berries (such as grapes, apples, or
peach The peach (''Prunus persica'') is a deciduous tree first domesticated and cultivated in Zhejiang province of Eastern China. It bears edible juicy fruits with various characteristics, most called peaches and others (the glossy-skinned, n ...
es), and exudates from plants (such as plant saps or cacti). Some yeasts are found in association with soil and insects. Yeasts from the soil and from the skins of fruits and berries have been shown to dominate fungal
succession Succession is the act or process of following in order or sequence. Governance and politics *Order of succession, in politics, the ascension to power by one ruler, official, or monarch after the death, resignation, or removal from office of ...
during fruit decay. The ecological function and
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...
of yeasts are relatively unknown compared to those of other
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in old ...
s. Yeasts, including '' Candida albicans'', '' Rhodotorula rubra'', '' Torulopsis'' and '' Trichosporon cutaneum'', have been found living in between people's toes as part of their skin flora. Yeasts are also present in the gut flora of mammals and some insects and even deep-sea environments host an array of yeasts. An Indian study of seven
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
species and nine plant species found 45 species from 16 genera colonize the nectaries of flowers and honey stomachs of bees. Most were members of the genus '' Candida''; the most common species in honey stomachs was '' Dekkera intermedia'' and in flower nectaries, '' Candida blankii''. Yeast colonising nectaries of the stinking hellebore have been found to raise the temperature of the flower, which may aid in attracting pollinators by increasing the evaporation of
volatile organic compound Volatile organic compounds (VOCs) are organic compounds that have a high vapour pressure at room temperature. High vapor pressure correlates with a low boiling point, which relates to the number of the sample's molecules in the surrounding air, a ...
s. A
black yeast “Black yeasts”, sometimes also black fungi, dematiaceous fungi, microcolonial fungi or meristematic fungi is a diverse group of slow-growing microfungi which reproduce mostly asexually (fungi imperfecti). Only few genera reproduce by budding ...
has been recorded as a partner in a complex relationship between ants, their mutualistic fungus, a fungal
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...
of the fungus and a bacterium that kills the parasite. The yeast has a negative effect on the bacteria that normally produce antibiotics to kill the parasite, so may affect the ants' health by allowing the parasite to spread. Certain strains of some species of yeasts produce proteins called yeast killer toxins that allow them to eliminate competing strains. (See main article on
killer yeast A killer yeast is a yeast, such as ''Saccharomyces cerevisiae'', which is able to secrete one of a number of toxic proteins which are lethal to susceptible cells. These "killer toxins" are polypeptides that kill sensitive cells of the same or relat ...
.) This can cause problems for winemaking but could potentially also be used to advantage by using killer toxin-producing strains to make the wine. Yeast killer toxins may also have medical applications in treating yeast infections (see "Pathogenic yeasts" section below). Marine yeasts, defined as the yeasts that are isolated from marine environments, are able to grow better on a medium prepared using seawater rather than freshwater. The first marine yeasts were isolated by Bernhard Fischer in 1894 from the Atlantic Ocean, and those were identified as ''Torula'' sp. and ''Mycoderma'' sp. Following this discovery, various other marine yeasts have been isolated from around the world from different sources, including seawater, seaweeds, marine fish and mammals. Among these isolates, some marine yeasts originated from terrestrial habitats (grouped as facultative marine yeast), which were brought to and survived in marine environments. The other marine yeasts were grouped as obligate or indigenous marine yeasts, which are confined to marine habitats. However, no sufficient evidence has been found to explain the indispensability of seawater for obligate marine yeasts. This article contains quotations from this source, which is available under a Creative Commons Attribution license. It has been reported that marine yeasts are able to produce many bioactive substances, such as amino acids, glucans, glutathione, toxins, enzymes, phytase, and vitamins with potential applications in the food, pharmaceutical, cosmetic, and chemical industries as well as for marine culture and environmental protection. Marine yeast was successfully used to produce bioethanol using seawater-based media which will potentially reduce the water footprint of bioethanol.


Reproduction

Yeasts, like all fungi, may have asexual and sexual reproductive cycles. The most common mode of vegetative growth in yeast is asexual reproduction by
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
, where a small bud (also known as a bleb or daughter cell) is formed on the parent cell. The
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: * Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...
of the parent cell splits into a daughter nucleus and migrates into the daughter cell. The bud then continues to grow until it separates from the parent cell, forming a new cell. The daughter cell produced during the budding process is generally smaller than the mother cell. Some yeasts, including '' Schizosaccharomyces pombe'', reproduce by fission instead of budding, and thereby creating two identically sized daughter cells. In general, under high-stress conditions such as
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excre ...
starvation, haploid cells will die; under the same conditions, however,
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectiv ...
cells can undergo sporulation, entering sexual reproduction (
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately ...
) and producing a variety of haploid
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s, which can go on to mate (conjugate), reforming the diploid. The haploid fission yeast ''Schizosaccharomyces pombe'' is a facultative sexual microorganism that can undergo mating when nutrients are limiting. Exposure of ''S. pombe'' to hydrogen peroxide, an agent that causes oxidative stress leading to oxidative DNA damage, strongly induces mating and the formation of meiotic spores. The budding yeast ''Saccharomyces cerevisiae'' reproduces by mitosis as diploid cells when nutrients are abundant, but when starved, this yeast undergoes meiosis to form haploid spores. Haploid cells may then reproduce asexually by mitosis. Katz Ezov et al. presented evidence that in natural ''S. cerevisiae'' populations clonal reproduction and selfing (in the form of intratetrad mating) predominate. In nature, mating of haploid cells to form diploid cells is most often between members of the same clonal population and
out-crossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a useful ...
is uncommon. Analysis of the ancestry of natural ''S. cerevisiae'' strains led to the conclusion that out-crossing occurs only about once every 50,000 cell divisions. These observations suggest that the possible long-term benefits of outcrossing (e.g. generation of diversity) are likely to be insufficient for generally maintaining sex from one generation to the next. Rather, a short-term benefit, such as recombinational repair during meiosis, may be the key to the maintenance of sex in ''S. cerevisiae''. Some pucciniomycete yeasts, in particular species of '' Sporidiobolus'' and ''
Sporobolomyces ''Sporobolomyces'' is a fungal genus of uncertain familial placement in the order Sporidiobolales. Species in the genus are anamorphic yeasts. Teleomorphic forms are included in the genus '' Sporidiobolus''. Generic synonyms of ''Sporobolomy ...
'', produce aerially dispersed, asexual ballistoconidia.


Uses

The useful physiological properties of yeast have led to their use in the field of
biotechnology Biotechnology is the integration of natural sciences and engineering sciences in order to achieve the application of organisms, cells, parts thereof and molecular analogues for products and services. The term ''biotechnology'' was first used ...
. Fermentation of sugars by yeast is the oldest and largest application of this technology. Many types of yeasts are used for making many foods: baker's yeast in bread production, brewer's yeast in beer fermentation, and yeast in wine fermentation and for xylitol production. So-called
red rice yeast Red yeast rice (), red rice ''koji'' (べにこうじ, lit. 'red ''koji), red fermented rice, red kojic rice, red ''koji'' rice, ''anka'', or ''angkak'', is a bright reddish purple fermented rice, which acquires its color from being cultivate ...
is actually a mold, '' Monascus purpureus''. Yeasts include some of the most widely used model organisms for
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar work ...
and cell biology.


Alcoholic beverages

Alcoholic beverages are defined as beverages that contain
ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a ...
(C2H5OH). This ethanol is almost always produced by fermentation – the
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run ...
of
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may o ...
s by certain species of yeasts under anaerobic or low-oxygen conditions. Beverages such as mead, wine, beer, or distilled spirits all use yeast at some stage of their production. A distilled beverage is a beverage containing ethanol that has been purified by
distillation Distillation, or classical distillation, is the process of separating the components or substances from a liquid mixture by using selective boiling and condensation, usually inside an apparatus known as a still. Dry distillation is the he ...
. Carbohydrate-containing plant material is fermented by yeast, producing a dilute solution of ethanol in the process. Spirits such as
whiskey Whisky or whiskey is a type of distilled alcoholic beverage made from fermented grain mash. Various grains (which may be malted) are used for different varieties, including barley, corn, rye, and wheat. Whisky is typically aged in wooden ...
and rum are prepared by distilling these dilute solutions of ethanol. Components other than ethanol are collected in the condensate, including water,
ester In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides ...
s, and other alcohols, which (in addition to that provided by the oak in which it may be aged) account for the flavour of the beverage.


Beer

Brewing yeasts may be classed as "top-cropping" (or "top-fermenting") and "bottom-cropping" (or "bottom-fermenting"). Top-cropping yeasts are so called because they form a foam at the top of the wort during fermentation. An example of a top-cropping yeast is ''
Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have b ...
'', sometimes called an "ale yeast". Bottom-cropping yeasts are typically used to produce lager-type beers, though they can also produce
ale Ale is a type of beer brewed using a warm fermentation method, resulting in a sweet, full-bodied and fruity taste. Historically, the term referred to a drink brewed without hops. As with most beers, ale typically has a bittering agent to bala ...
-type beers. These yeasts ferment well at low temperatures. An example of bottom-cropping yeast is '' Saccharomyces pastorianus'', formerly known as ''S. carlsbergensis''. Decades ago, taxonomists reclassified ''S. carlsbergensis'' (uvarum) as a member of ''S. cerevisiae'', noting that the only distinct difference between the two is metabolic. Lager strains of ''S. cerevisiae'' secrete an enzyme called melibiase, allowing them to hydrolyse melibiose, a disaccharide, into more fermentable
monosaccharide Monosaccharides (from Greek '' monos'': single, '' sacchar'': sugar), also called simple sugars, are the simplest forms of sugar and the most basic units (monomers) from which all carbohydrates are built. They are usually colorless, water- so ...
s. Top- and bottom-cropping and cold- and warm-fermenting distinctions are largely generalizations used by laypersons to communicate to the general public. The most common top-cropping brewer's yeast, ''S. cerevisiae'', is the same species as the common baking yeast. Brewer's yeast is also very rich in
essential mineral In the context of nutrition, a mineral is a chemical element required as an essential nutrient by organisms to perform functions necessary for life. However, the four major structural elements in the human body by weight (oxygen, hydrogen, carbon ...
s and the B vitamins (except B12), a feature exploited in food products made from leftover (
by-product A by-product or byproduct is a secondary product derived from a production process, manufacturing process or chemical reaction; it is not the primary product or service being produced. A by-product can be useful and marketable or it can be consid ...
) yeast from brewing. However, baking and brewing yeasts typically belong to different strains, cultivated to favour different characteristics: baking yeast strains are more aggressive, to carbonate
dough Dough is a thick, malleable, sometimes elastic paste made from grains or from leguminous or chestnut crops. Dough is typically made by mixing flour with a small amount of water or other liquid and sometimes includes yeast or other leavenin ...
in the shortest amount of time possible; brewing yeast strains act more slowly but tend to produce fewer off-flavours and tolerate higher alcohol concentrations (with some strains, up to 22%). '' Dekkera/Brettanomyces'' is a genus of yeast known for its important role in the production of '
lambic Lambic () is a type of beer brewed in the Pajottenland region of Belgium southwest of Brussels and in Brussels itself since the 13th century. Types of lambic beers include gueuze, kriek lambic and framboise. Lambic differs from most other be ...
' and specialty
sour ale Sour beer, also known as Sours, is beer which has an intentionally acidic, tart, or sour taste. Traditional sour beer styles include Belgian lambics, gueuze and Flanders red ale, and German gose and Berliner Weisse. Brewing Unlike modern brewin ...
s, along with the secondary conditioning of a particular Belgian Trappist beer. The taxonomy of the genus ''Brettanomyces'' has been debated since its early discovery and has seen many reclassifications over the years. Early classification was based on a few species that reproduced asexually (anamorph form) through multipolar budding. Shortly after, the formation of ascospores was observed and the genus ''Dekkera'', which reproduces sexually (teleomorph form), was introduced as part of the taxonomy. The current taxonomy includes five species within the genera of ''Dekkera/Brettanomyces''. Those are the anamorphs '' Brettanomyces bruxellensis'', ''
Brettanomyces anomalus ''Brettanomyces'' is a non-spore forming genus of yeast in the family Saccharomycetaceae, and is often colloquially referred to as "Brett". The genus name ''Dekkera'' is used interchangeably with ''Brettanomyces'', as it describes the teleomorph ...
'', '' Brettanomyces custersianus'', '' Brettanomyces naardenensis'', and ''
Brettanomyces nanus ''Brettanomyces'' is a spore, non-spore forming genus of yeast in the family Saccharomycetaceae, and is often colloquially referred to as "Brett". The genus name ''Dekkera'' is used interchangeably with ''Brettanomyces'', as it describes the tele ...
'', with teleomorphs existing for the first two species, ''
Dekkera bruxellensis ''Brettanomyces bruxellensis'' (the anamorph of ''Dekkera bruxellensis'') is a yeast associated with the Senne valley near Brussels, Belgium. Despite its Latin species name, ''B. bruxellensis'' is found all over the globe. In the wild, it is oft ...
'' and ''
Dekkera anomala ''Brettanomyces'' is a spore, non-spore forming genus of yeast in the family Saccharomycetaceae, and is often colloquially referred to as "Brett". The genus name ''Dekkera'' is used interchangeably with ''Brettanomyces'', as it describes the tele ...
''. The distinction between ''Dekkera'' and ''Brettanomyces'' is arguable, with Oelofse et al. (2008) citing Loureiro and Malfeito-Ferreira from 2006 when they affirmed that current molecular DNA detection techniques have uncovered no variance between the anamorph and teleomorph states. Over the past decade, ''Brettanomyces'' spp. have seen an increasing use in the craft-brewing sector of the industry, with a handful of breweries having produced beers that were primarily fermented with pure cultures of ''Brettanomyces'' spp. This has occurred out of experimentation, as very little information exists regarding pure culture fermentative capabilities and the aromatic compounds produced by various strains. ''Dekkera''/''Brettanomyces'' spp. have been the subjects of numerous studies conducted over the past century, although a majority of the recent research has focused on enhancing the knowledge of the wine industry. Recent research on eight ''Brettanomyces'' strains available in the brewing industry focused on strain-specific fermentations and identified the major compounds produced during pure culture anaerobic fermentation in wort.


Wine

Yeast is used in
winemaking Winemaking or vinification is the production of wine, starting with the selection of the fruit, its fermentation into alcohol, and the bottling of the finished liquid. The history of wine-making stretches over millennia. The science of wine and ...
, where it converts the sugars present (
glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...
and fructose) in
grape juice Grape juice is obtained from crushing and blending grapes into a liquid. In the wine industry, grape juice that contains 7–23 percent of pulp, skins, stems and seeds is often referred to as '' must''. The sugars in grape juice allow it to ...
( must) into ethanol. Yeast is normally already present on grape skins. Fermentation can be done with this endogenous "wild yeast", but this procedure gives unpredictable results, which depend upon the exact types of yeast species present. For this reason, a pure yeast culture is usually added to the must; this yeast quickly dominates the fermentation. The wild yeasts are repressed, which ensures a reliable and predictable fermentation. Most added wine yeasts are strains of ''S. cerevisiae'', though not all strains of the species are suitable. Different ''S. cerevisiae'' yeast strains have differing physiological and fermentative properties, therefore the actual strain of yeast selected can have a direct impact on the finished wine. Significant research has been undertaken into the development of novel wine yeast strains that produce atypical flavour profiles or increased complexity in wines. The growth of some yeasts, such as '' Zygosaccharomyces'' and '' Brettanomyces'', in wine can result in wine faults and subsequent spoilage. ''Brettanomyces'' produces an array of
metabolite In biochemistry, a metabolite is an intermediate or end product of metabolism. The term is usually used for small molecules. Metabolites have various functions, including fuel, structure, signaling, stimulatory and inhibitory effects on enzymes, ...
s when growing in wine, some of which are volatile
phenol Phenol (also called carbolic acid) is an aromatic organic compound with the molecular formula . It is a white crystalline solid that is volatile. The molecule consists of a phenyl group () bonded to a hydroxy group (). Mildly acidic, it r ...
ic compounds. Together, these compounds are often referred to as "''Brettanomyces'' character", and are often described as " antiseptic" or "barnyard" type aromas. ''Brettanomyces'' is a significant contributor to wine faults within the wine industry. Researchers from the
University of British Columbia The University of British Columbia (UBC) is a public research university with campuses near Vancouver and in Kelowna, British Columbia. Established in 1908, it is British Columbia's oldest university. The university ranks among the top thr ...
, Canada, have found a new strain of yeast that has reduced
amine In chemistry, amines (, ) are compounds and functional groups that contain a basic nitrogen atom with a lone pair. Amines are formally derivatives of ammonia (), wherein one or more hydrogen atoms have been replaced by a substituent ...
s. The amines in red wine and
Chardonnay Chardonnay (, , ) is a green-skinned grape variety used in the production of white wine. The variety originated in the Burgundy wine region of eastern France, but is now grown wherever wine is produced, from England to New Zealand. For new ...
produce off-flavors and cause headaches and hypertension in some people. About 30% of people are sensitive to biogenic amines, such as
histamine Histamine is an organic nitrogenous compound involved in local immune responses, as well as regulating physiological functions in the gut and acting as a neurotransmitter for the brain, spinal cord, and uterus. Since histamine was discover ...
s.


Baking

Yeast, the most common one being ''S. cerevisiae'', is used in baking as a leavening agent, where it converts the
food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is in ...
/fermentable sugars present in dough into the gas
carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...
. This causes the dough to expand or rise as gas forms pockets or bubbles. When the dough is baked, the yeast dies and the air pockets "set", giving the baked product a soft and spongy texture. The use of potatoes, water from potato boiling, eggs, or sugar in a bread dough accelerates the growth of yeasts. Most yeasts used in baking are of the same species common in alcoholic fermentation. In addition, ''Saccharomyces exiguus'' (also known as ''S. minor''), a wild yeast found on plants, fruits, and grains, is occasionally used for baking. In breadmaking, the yeast initially respires aerobically, producing carbon dioxide and water. When the oxygen is depleted, fermentation begins, producing ethanol as a waste product; however, this evaporates during baking. It is not known when yeast was first used to bake bread. The first records that show this use came from Ancient Egypt. Researchers speculate a mixture of flour meal and water was left longer than usual on a warm day and the yeasts that occur in natural contaminants of the
flour Flour is a powder made by grinding raw grains, roots, beans, nuts, or seeds. Flours are used to make many different foods. Cereal flour, particularly wheat flour, is the main ingredient of bread, which is a staple food for many cul ...
caused it to ferment before baking. The resulting bread would have been lighter and tastier than the normal flat, hard cake. Today, there are several retailers of baker's yeast; one of the earlier developments in North America is Fleischmann's Yeast, in 1868. During World War II, Fleischmann's developed a
granulated Granulation is the process of forming grains or granules from a powdery or solid substance, producing a granular material. It is applied in several technological processes in the chemical and pharmaceutical industries. Typically, granulation invo ...
active dry yeast which did not require refrigeration, had a longer shelf life than fresh yeast, and rose twice as fast. Baker's yeast is also sold as a fresh yeast compressed into a square "cake". This form perishes quickly, so must be used soon after production. A weak solution of water and sugar can be used to determine whether yeast is expired. In the solution, active yeast will foam and bubble as it ferments the sugar into ethanol and carbon dioxide. Some recipes refer to this as proofing the yeast, as it "proves" (tests) the viability of the yeast before the other ingredients are added. When a sourdough starter is used, flour and water are added instead of sugar; this is referred to as proofing the
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...
. When yeast is used for making bread, it is mixed with
flour Flour is a powder made by grinding raw grains, roots, beans, nuts, or seeds. Flours are used to make many different foods. Cereal flour, particularly wheat flour, is the main ingredient of bread, which is a staple food for many cul ...
, salt, and warm water or milk. The dough is kneaded until it is smooth, and then left to rise, sometimes until it has doubled in size. The dough is then shaped into loaves. Some bread doughs are knocked back after one rising and left to rise again (this is called dough proofing) and then baked. A longer rising time gives a better flavor, but the yeast can fail to raise the bread in the final stages if it is left for too long initially.


Bioremediation

Some yeasts can find potential application in the field of bioremediation. One such yeast, ''
Yarrowia lipolytica ''Yarrowia'' is a fungal genus in the family Dipodascaceae. For a while the genus was monotypic, containing the single species ''Yarrowia lipolytica'', a yeast that can use unusual carbon sources, such as hydrocarbons. This has made it of interes ...
'', is known to degrade palm oil mill effluent, TNT (an explosive material), and other
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ...
s, such as
alkane In organic chemistry, an alkane, or paraffin (a historical trivial name that also has other meanings), is an acyclic saturated hydrocarbon. In other words, an alkane consists of hydrogen and carbon atoms arranged in a tree structure in wh ...
s,
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, f ...
s, fats and oils. It can also tolerate high concentrations of salt and
heavy metals upright=1.2, Crystals of osmium, a heavy metal nearly twice as dense as lead">lead.html" ;"title="osmium, a heavy metal nearly twice as dense as lead">osmium, a heavy metal nearly twice as dense as lead Heavy metals are generally defined as ...
, and is being investigated for its potential as a heavy metal biosorbent. ''Saccharomyces cerevisiae'' has potential to bioremediate toxic pollutants like
arsenic Arsenic is a chemical element with the symbol As and atomic number 33. Arsenic occurs in many minerals, usually in combination with sulfur and metals, but also as a pure elemental crystal. Arsenic is a metalloid. It has various allotropes, b ...
from industrial effluent. Bronze statues are known to be degraded by certain species of yeast. Different yeasts from Brazilian gold mines bioaccumulate free and complexed silver ions.


Industrial ethanol production

The ability of yeast to convert sugar into ethanol has been harnessed by the biotechnology industry to produce
ethanol fuel Ethanol fuel is ethyl alcohol, the same type of alcohol found in alcoholic beverages, used as fuel. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline. The first production car running entirely on ethanol was t ...
. The process starts by milling a feedstock, such as sugar cane,
field corn Field corn, also known as cow corn, is a North American term for maize (''Zea mays'') grown for livestock fodder (silage), ethanol, cereal, and processed food products. The principal field corn varieties are dent corn, flint corn, flour corn (a ...
, or other
cereal grain A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
s, and then adding dilute
sulfuric acid Sulfuric acid (American spelling and the preferred IUPAC name) or sulphuric acid ( Commonwealth spelling), known in antiquity as oil of vitriol, is a mineral acid composed of the elements sulfur, oxygen and hydrogen, with the molecular fo ...
, or fungal alpha amylase enzymes, to break down the starches into complex sugars. A glucoamylase is then added to break the complex sugars down into simple sugars. After this, yeasts are added to convert the simple sugars to ethanol, which is then distilled off to obtain ethanol up to 96% in purity. ''Saccharomyces'' yeasts have been genetically engineered to ferment
xylose Xylose ( grc, ξύλον, , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional g ...
, one of the major fermentable sugars present in cellulosic biomasses, such as agriculture residues, paper wastes, and wood chips. Such a development means ethanol can be efficiently produced from more inexpensive feedstocks, making cellulosic ethanol fuel a more competitively priced alternative to gasoline fuels.


Nonalcoholic beverages

A number of sweet carbonated beverages can be produced by the same methods as beer, except the fermentation is stopped sooner, producing carbon dioxide, but only trace amounts of alcohol, leaving a significant amount of residual sugar in the drink. * Root beer, originally made by Native Americans, commercialized in the United States by Charles Elmer Hires and especially popular during
Prohibition Prohibition is the act or practice of forbidding something by law; more particularly the term refers to the banning of the manufacture, storage (whether in barrels or in bottles), transportation, sale, possession, and consumption of alcoholi ...
* Kvass, a fermented drink made from rye, popular in Eastern Europe. It has a recognizable, but low alcoholic content. * Kombucha, a fermented sweetened tea. Yeast in
symbiosis Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...
with acetic acid bacteria is used in its preparation. Species of yeasts found in the tea can vary, and may include: '' Brettanomyces bruxellensis'', '' Candida stellata'', '' Schizosaccharomyces pombe'', ''
Torulaspora delbrueckii ''Torulaspora delbrueckii'' is a ubiquitous yeast species with both wild and anthropic habitats. The type strain of ''T. delbrueckii'' is CBS 1146T, equivalent to CLIB 230 or ATCC 10662etc. The type strain of ''T. delbrueckii'' CBS 1146 T was seq ...
'' and ''
Zygosaccharomyces bailii ''Zygosaccharomyces bailii'' is a species in the genus '' Zygosaccharomyces''. It was initially described as ''Saccharomyces bailii'' by Lindner in 1895, but in 1983 it was reclassified as ''Zygosaccharomyces bailii'' in the work by Barnett et a ...
''. Also popular in Eastern Europe and some former Soviet republics under the name ''chajnyj grib'' (russian: Чайный гриб), which means "tea mushroom". * Kefir and
kumis ''Kumis'' (also spelled ''kumiss'' or ''koumiss'' or ''kumys'', see other transliterations and cognate words below under terminology and etymology – otk, airag kk, қымыз, ''qymyz'') mn, айраг, ''ääryg'') is a fermented dairy ...
are made by fermenting milk with yeast and bacteria. * Mauby ( es, mabí), made by fermenting sugar with the wild yeasts naturally present on the bark of the '' Colubrina elliptica'' tree, popular in the
Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean ...


Foods and

Yeast is used as an ingredient in foods for its umami flavor, in much of the same way that
monosodium glutamate Monosodium glutamate (MSG), also known as sodium glutamate, is the sodium salt of glutamic acid. MSG is found naturally in some foods including tomatoes and cheese in this glutamic acid form. MSG is used in cooking as a flavor enhancer ...
(MSG) is used and, like MSG, often contain free glutamic acid. Examples include: * Yeast extract, made from the intracellular contents of yeast and used as food additives or flavours. The general method for making yeast extract for food products such as Vegemite and Marmite on a commercial scale is ''heat autolysis'', i.e. to add salt to a suspension of yeast, making the solution hypertonic, which leads to the cells' shrivelling up. This triggers autolysis, wherein the yeast's digestive enzymes break their own
protein Proteins are large biomolecules and macromolecules that comprise one or more long chains of amino acid residues. Proteins perform a vast array of functions within organisms, including catalysing metabolic reactions, DNA replication, res ...
s down into simpler compounds, a process of self-destruction. The dying yeast cells are then heated to complete their breakdown, after which the husks (yeast with thick cell walls that would give poor texture) are removed. Yeast autolysates are used in Vegemite and Promite (Australia); Marmite (the United Kingdom); the unrelated Marmite (New Zealand); Vitam-R (Germany); and Cenovis (
Switzerland ). Swiss law does not designate a ''capital'' as such, but the federal parliament and government are installed in Bern, while other federal institutions, such as the federal courts, are in other cities (Bellinzona, Lausanne, Luzern, Neuchâtel ...
). * Nutritional yeast, which is whole dried, deactivated yeast cells, usually ''S. cerevisiae''. Usually in the form of yellow flake or powder, its nutty and umami flavor makes it a vegan substitute for cheese powder. Another popular use is as a topping for popcorn. It can also be used in mashed and fried potatoes, as well as in
scrambled egg Scrambled eggs is a dish made from eggs (usually chicken eggs) stirred, whipped or beaten together while being gently heated, typically with salt, butter, oil and sometimes other ingredients. Preparation Only eggs are necessary to make scramble ...
s. It comes in the form of flakes, or as a yellow powder similar in texture to cornmeal. In Australia, it is sometimes sold as "savoury yeast flakes". Both types of yeast foods above are rich in B-complex vitamins (besides vitamin B12 unless fortified), making them an attractive nutritional supplement to vegans. The same vitamins are also found in some yeast-fermented products mentioned above, such as kvass. Nutritional yeast in particular is naturally low in fat and
sodium Sodium is a chemical element with the symbol Na (from Latin ''natrium'') and atomic number 11. It is a soft, silvery-white, highly reactive metal. Sodium is an alkali metal, being in group 1 of the periodic table. Its only stable ...
and a source of protein and vitamins as well as other minerals and
cofactors Cofactor may also refer to: * Cofactor (biochemistry), a substance that needs to be present in addition to an enzyme for a certain reaction to be catalysed * A domain parameter in elliptic curve cryptography, defined as the ratio between the order ...
required for growth. Many brands of nutritional yeast and yeast extract spreads, though not all, are fortified with vitamin B12, which is produced separately by
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
. In 1920, the Fleischmann Yeast Company began to promote yeast cakes in a "Yeast for Health" campaign. They initially emphasized yeast as a source of vitamins, good for skin and digestion. Their later advertising claimed a much broader range of health benefits, and was censured as misleading by the
Federal Trade Commission The Federal Trade Commission (FTC) is an independent agency of the United States government whose principal mission is the enforcement of civil (non-criminal) antitrust law and the promotion of consumer protection. The FTC shares jurisdiction o ...
. The fad for yeast cakes lasted until the late 1930s.


Probiotics

Some
probiotic Probiotics are live microorganisms promoted with claims that they provide health benefits when consumed, generally by improving or restoring the gut microbiota. Probiotics are considered generally safe to consume, but may cause bacteria-host i ...
supplements use the yeast '' S. boulardii'' to maintain and restore the natural flora in the gastrointestinal tract. ''S. boulardii'' has been shown to reduce the symptoms of acute
diarrhea Diarrhea, also spelled diarrhoea, is the condition of having at least three loose, liquid, or watery bowel movements each day. It often lasts for a few days and can result in dehydration due to fluid loss. Signs of dehydration often begin w ...
, reduce the chance of infection by '' Clostridium difficile'' (often identified simply as C. difficile or C. diff), reduce bowel movements in diarrhea-predominant IBS patients, and reduce the incidence of
antibiotic An antibiotic is a type of antimicrobial substance active against bacteria. It is the most important type of antibacterial agent for fighting bacterial infections, and antibiotic medications are widely used in the treatment and prevention ...
-, traveler's-, and
HIV/AIDS Human immunodeficiency virus infection and acquired immunodeficiency syndrome (HIV/AIDS) is a spectrum of conditions caused by infection with the human immunodeficiency virus (HIV), a retrovirus. Following initial infection an individual ...
-associated diarrheas.


Aquarium hobby

Yeast is often used by
aquarium An aquarium (plural: ''aquariums'' or ''aquaria'') is a vivarium of any size having at least one transparent side in which aquatic plants or animals are kept and displayed. Fishkeepers use aquaria to keep fish, invertebrates, amphibians, aq ...
hobbyists to generate carbon dioxide (CO2) to nourish plants in planted aquaria. CO2 levels from yeast are more difficult to regulate than those from pressurized CO2 systems. However, the low cost of yeast makes it a widely used alternative.


Scientific research

Several yeasts, in particular '' S. cerevisiae'' and '' S. pombe'', have been widely used in genetics and cell biology, largely because they are simple
eukaryotic Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacte ...
cells, serving as a model for all eukaryotes, including humans, for the study of fundamental cellular processes such as the
cell cycle The cell cycle, or cell-division cycle, is the series of events that take place in a cell that cause it to divide into two daughter cells. These events include the duplication of its DNA (DNA replication) and some of its organelles, and sub ...
,
DNA replication In molecular biology, DNA replication is the biological process of producing two identical replicas of DNA from one original DNA molecule. DNA replication occurs in all living organisms acting as the most essential part for biological inheritan ...
, recombination,
cell division Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ...
, and metabolism. Also, yeasts are easily manipulated and cultured in the laboratory, which has allowed for the development of powerful standard techniques, such as yeast two-hybrid,
synthetic genetic array Synthetic genetic array analysis (SGA) is a high-throughput technique for exploring synthetic lethal and synthetic sick genetic interactions (SSL). SGA allows for the systematic construction of double mutants using a combination of recombinant g ...
analysis, and tetrad analysis. Many proteins important in human biology were first discovered by studying their homologues in yeast; these proteins include cell cycle proteins, signaling proteins, and protein-processing
enzyme Enzymes () are proteins that act as biological catalysts by accelerating chemical reactions. The molecules upon which enzymes may act are called substrates, and the enzyme converts the substrates into different molecules known as products ...
s. On 24 April 1996, ''S. cerevisiae'' was announced to be the first eukaryote to have its
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ...
, consisting of 12 million
base pair A base pair (bp) is a fundamental unit of double-stranded nucleic acids consisting of two nucleobases bound to each other by hydrogen bonds. They form the building blocks of the DNA double helix and contribute to the folded structure of both D ...
s, fully sequenced as part of the
Genome Project Genome projects are scientific endeavours that ultimately aim to determine the complete genome sequence of an organism (be it an animal, a plant, a fungus, a bacterium, an archaean, a protist or a virus) and to annotate protein-coding genes ...
. At the time, it was the most complex organism to have its full genome sequenced, and the work seven years and the involvement of more than 100 laboratories to accomplish. The second yeast species to have its genome sequenced was ''Schizosaccharomyces pombe'', which was completed in 2002. It was the sixth eukaryotic genome sequenced and consists of 13.8 million base pairs. As of 2014, over 50 yeast species have had their genomes sequenced and published. Genomic and functional gene annotation of the two major yeast models can be accessed via their respective model organism databases: SGD and PomBase.


Genetically engineered biofactories

Various yeast species have been genetically engineered to efficiently produce various drugs, a technique called
metabolic engineering Metabolic engineering is the practice of optimizing genetic and regulatory processes within cells to increase the cell's production of a certain substance. These processes are chemical networks that use a series of biochemical reactions and enzym ...
. ''S. cerevisiae'' is easy to genetically engineer; its physiology, metabolism and genetics are well known, and it is amenable for use in harsh industrial conditions. A wide variety of chemical in different classes can be produced by engineered yeast, including phenolics, isoprenoids,
alkaloid Alkaloids are a class of basic, naturally occurring organic compounds that contain at least one nitrogen atom. This group also includes some related compounds with neutral and even weakly acidic properties. Some synthetic compounds of simila ...
s, and polyketides. About 20% of biopharmaceuticals are produced in ''S. cerevisiae'', including
insulin Insulin (, from Latin ''insula'', 'island') is a peptide hormone produced by beta cells of the pancreatic islets encoded in humans by the ''INS'' gene. It is considered to be the main anabolic hormone of the body. It regulates the metabolism ...
,
vaccine A vaccine is a biological preparation that provides active acquired immunity to a particular infectious or malignant disease. The safety and effectiveness of vaccines has been widely studied and verified.
s for
hepatitis Hepatitis is inflammation of the liver tissue. Some people or animals with hepatitis have no symptoms, whereas others develop yellow discoloration of the skin and whites of the eyes ( jaundice), poor appetite, vomiting, tiredness, abdominal ...
, and human serum albumin.


Pathogenic yeasts

Some species of yeast are opportunistic pathogens that can cause infection in people with compromised
immune system The immune system is a network of biological processes that protects an organism from diseases. It detects and responds to a wide variety of pathogens, from viruses to parasitic worms, as well as cancer cells and objects such as wood splinte ...
s. ''
Cryptococcus neoformans ''Cryptococcus neoformans'' is an encapsulated yeast belonging to the class Tremellomycetes and an obligate aerobe that can live in both plants and animals. Its teleomorph is a filamentous fungus, formerly referred to ''Filobasidiella neoformans ...
'' and '' Cryptococcus gattii'' are significant pathogens of immunocompromised people. They are the species primarily responsible for
cryptococcosis Cryptococcosis is a potentially fatal fungal infection of mainly the lungs, presenting as a pneumonia, and brain, where it appears as a meningitis. Cough, difficulty breathing, chest pain and fever are seen when the lungs are infected. When the ...
, a fungal infection that occurs in about one million
HIV/AIDS Human immunodeficiency virus infection and acquired immunodeficiency syndrome (HIV/AIDS) is a spectrum of conditions caused by infection with the human immunodeficiency virus (HIV), a retrovirus. Following initial infection an individual ...
patients, causing over 600,000 deaths annually. The cells of these yeast are surrounded by a rigid
polysaccharide Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with w ...
capsule, which helps to prevent them from being recognised and engulfed by white blood cells in the human body. Yeasts of the genus '' Candida'', another group of opportunistic pathogens, cause
oral The word oral may refer to: Relating to the mouth * Relating to the mouth, the first portion of the alimentary canal that primarily receives food and liquid ** Oral administration of medicines ** Oral examination (also known as an oral exam or or ...
and vaginal infections in humans, known as candidiasis. ''Candida'' is commonly found as a commensal yeast in the mucous membranes of humans and other warm-blooded animals. However, sometimes these same strains can become pathogenic. The yeast cells sprout a
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
l outgrowth, which locally penetrates the
mucosal membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It is ...
, causing irritation and shedding of the tissues. A book from the 1980s listed the pathogenic yeasts of candidiasis in probable descending order of
virulence Virulence is a pathogen's or microorganism's ability to cause damage to a host. In most, especially in animal systems, virulence refers to the degree of damage caused by a microbe to its host. The pathogenicity of an organism—its ability to ...
for humans as: '' C. albicans'', '' C. tropicalis'', '' C. stellatoidea'', '' C. glabrata'', '' C. krusei'', '' C. parapsilosis'', '' C. guilliermondii'', '' C. viswanathii'', '' C. lusitaniae'', and ''
Rhodotorula mucilaginosa ''Rhodotorula'' is a genus of pigmented yeasts, part of the division Basidiomycota. It is readily identifiable by distinctive orange/red colonies when grown on Sabouraud's dextrose agar (SDA). This distinctive color is the result of pigments ...
''. ''Candida glabrata'' is the second most common ''Candida'' pathogen after ''C. albicans'', causing infections of the
urogenital tract The genitourinary system, or urogenital system, are the organs of the reproductive system and the urinary system. These are grouped together because of their proximity to each other, their common embryological origin and the use of common pathwa ...
, and of the bloodstream (
candidemia Fungemia is the presence of fungi or yeasts in the blood. The most common type, also known as candidemia, candedemia, or systemic candidiasis, is caused by '' Candida'' species; candidemia is also among the most common bloodstream infections of ...
). '' C. auris'' has been more recently identified.


Food spoilage

Yeasts are able to grow in foods with a low pH (5.0 or lower) and in the presence of sugars, organic acids, and other easily metabolized carbon sources. During their growth, yeasts metabolize some food components and produce metabolic end products. This causes the physical, chemical, and sensible properties of a food to change, and the food is spoiled. The growth of yeast within food products is often seen on their surfaces, as in cheeses or meats, or by the fermentation of sugars in beverages, such as juices, and semiliquid products, such as syrups and jams. The yeast of the genus '' Zygosaccharomyces'' have had a long history as spoilage yeasts within the food industry. This is mainly because these species can grow in the presence of high sucrose, ethanol, acetic acid, sorbic acid,
benzoic acid Benzoic acid is a white (or colorless) solid organic compound with the formula , whose structure consists of a benzene ring () with a carboxyl () substituent. It is the simplest aromatic carboxylic acid. The name is derived from gum benzoin ...
, and
sulfur dioxide Sulfur dioxide (IUPAC-recommended spelling) or sulphur dioxide (traditional Commonwealth English) is the chemical compound with the formula . It is a toxic gas responsible for the odor of burnt matches. It is released naturally by volcanic a ...
concentrations, representing some of the commonly used
food preservation Food preservation includes processes that make food more resistant to microorganism growth and slow the oxidation of fats. This slows down the decomposition and rancidification process. Food preservation may also include processes that inhibit ...
methods.
Methylene blue Methylthioninium chloride, commonly called methylene blue, is a salt used as a dye and as a medication. Methylene blue is a thiazine dye. As a medication, it is mainly used to treat methemoglobinemia by converting the ferric iron in hemoglobin ...
is used to test for the presence of live yeast cells. In oenology, the major spoilage yeast is '' Brettanomyces bruxellensis''. '' Candida blankii'' has been detected in Iberian ham and meat.


Symbiosis

An Indian study of seven
bee Bees are winged insects closely related to wasps and ants, known for their roles in pollination and, in the case of the best-known bee species, the western honey bee, for producing honey. Bees are a monophyletic lineage within the superfami ...
species and nine plant species found 45 yeast species from 16 genera colonise the nectaries of flowers and honey stomachs of bees. Most were members of the genus '' Candida''; the most common species in honey bee stomachs was '' Dekkera intermedia'', while the most common species colonising flower nectaries was '' Candida blankii''. Although the mechanics are not fully understood, it was found that ''A. indica'' flowers more if '' Candida blankii'' is present. In another example, ''Spathaspora passalidarum'', found in the digestive tract of scarab beetles, aids the digestion of plant cells by fermenting
xylose Xylose ( grc, ξύλον, , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional g ...
.


See also

*
Bioaerosol Bioaerosols (short for biological aerosols) are a subcategory of particles released from terrestrial and marine ecosystems into the atmosphere. They consist of both living and non-living components, such as fungi, pollen, bacteria and viruses. Commo ...
* Ethanol fermentation *
Evolution of aerobic fermentation Aerobic fermentation or aerobic glycolysis is a metabolic process by which cells metabolize sugars via fermentation in the presence of oxygen and occurs through the repression of normal respiratory metabolism. It is referred to as the Crabtree effe ...
* '' Kazachstania yasuniensis'' – a yeast isolated in 2015 * Mycosis (fungal infection in animals) *
Start point (yeast) The Start checkpoint is a major cell cycle checkpoint in yeast. The Start checkpoint ensures irreversible cell-cycle entry even if conditions later become unfavorable. The physiological factors that control passage through the Start checkpoint incl ...
* WHI3 * Yeast plasmids * Zymology


References


Further reading

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External links


''Saccharomyces'' genome database

Yeast growth and the cell cycle

Yeast virtual library
{{Authority control Abortifacients Brewing ingredients Fermentation in food processing Food additives Fungus common names Leavening agents Medicinal fungi